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Ecophysiological aspects of guava (Psidium guajava L.). A review - Revistas UPTC
Doi: https://doi.org/10.17584/rcch.2021v15i2.12355

Ecophysiological aspects of guava (Psidium guajava L.).
A review
Aspectos de la ecofisiología de la guayaba (Psidium guajava L.).
Una revisión

                                                                   GERHARD FISCHER1, 3
                                                                   LUZ MARINA MELGAREJO2

                                                                Fruiting guava plant.
                                                                Photo: G. Fischer

ABSTRACT
Guava, because of its ability to grow in tropical and subtropical climates, has been introduced to some 60
low-latitude countries. It is adapted to a temperature range between 15 and 30°C. Outside this range, the
effect of lower or higher temperatures reduces fruit set, while night temperatures of 5 to 7°C stop growth.
Additionally, low temperatures hinder production, causing flowers to fall or increasing the fruit development
cycle, up to 220 days. When estimating the cardinal temperatures of development, the minimum threshold
temperature was 10.9°C, the optimum temperature was 17.3°C, and the maximum threshold temperature
was 51.2°C. The guava tree adapts well to altitudes between 0 and 2,000 m a.s.l. in Colombia; however, there
is a high genotype×environment interaction for production and quality characteristics in fruits with respect
to the orchard elevation. Radiation >2,000 µmol photons m-2 s-1 decreased the fruit ascorbic acid content. An
important ecophysiological factor in guava is water, and crops require between 1,000 to 2,000 mm year-1. It
withstands waterlogging for several days, but excess precipitation and atmospheric humidity decrease fruit
quality considerably. However, this tree is classified as moderately drought-tolerant to stress from water
deficits, affecting flowering and fruit set. It is also moderately tolerant to salinity, depending on the variety,
supporting electrical conductivities up to 1.5-1.8 dS m-1. Generally, guava can be cultivated in a wide range of
tropical and subtropical areas, where it is preferred because of its high nutritional and medicinal contents and
its aptitude for transport and handling.

                                               Additional keywords: temperature; altitude; humidity; fruit quality; salinity.

1
    Independent Consulter, Emeritus Researcher of Colciencias, Bogota (Colombia). ORCID Fischer, G.:
    0000-0001-8101-0507
2
    Universidad Nacional de Colombia, Faculty of Sciences, Department of Biology, Bogota (Colombia).
    ORCID Melgarejo, L.M.: 0000-0003-3148-1911
3
    Corresponding author. gerfischer@gmail.com

                                         REVISTA COLOMBIANA DE CIENCIAS HORTÍCOLAS - Vol. 15 - No. 2, e12355, May-August 2021
                                                                                       e-ISSN: 2422-3719 · ISSN-L: 2011-2173
2              FISCHER / MELGAREJO

RESUMEN
La guayaba, debido a su habilidad de crecer en climas tropicales y subtropicales, ha sido introducida en unos 60
países de las latitudes bajas. Se adapta a rangos de temperatura entre los 15 y 30°C. Fuera de este rango, el efec-
to de temperaturas inferiores o superiores reducen, en primer lugar, el cuajamiento de los frutos, y temperaturas
nocturnas de 5 a 7°C detienen el crecimiento. Adicionalmente, las temperaturas bajas dificultan la producción
generando caída de flores o aumentando el ciclo de desarrollo del fruto hasta unos 220 días. En una estimación de
las temperaturas cardinales de desarrollo se encontraron como temperatura umbral mínima 10,9°C, temperatura
óptima 17,3°C y temperatura umbral máxima 51,2°C. El árbol de guayaba se adapta bien a altitudes entre 0 y 2.000
msnm en Colombia; sin embargo, existe alta interacción genotipo (variedad)×ambiente referente a las caracterís-
ticas de producción y calidad del fruto con respecto a la elevación del sitio. Radiaciones >2.000 µmol fotones m-2
s-1 disminuyeron el contenido del ácido ascórbico en el fruto. Un factor ecofisiológico importante en la guayaba es
el agua ya que los cultivos exigen entre unos 1.000 a 2.000 mm año -1. Soportan el anegamiento de varios días; pero
mucha precipitación y humedad atmosférica disminuyen la calidad del fruto considerablemente. No obstante, este
árbol está clasificado como moderadamente tolerante a la sequía, el estrés por déficit hídrico afecta la floración y
el cuajamiento de los frutos. Es también moderadamente tolerante a la salinidad, dependiendo de la variedad, con
conductividades eléctricas hasta máximo 1,5-1,8 dS m-1. En general, se puede concluir que la guayaba se puede cul-
tivar en una amplia gama de áreas tropicales y subtropicales donde es preferida por su alto contenido nutricional y
medicinal y su aptitud para el transporte y manejo.

Palabras clave adicionales: temperatura; altitud; humedad; calidad fruto; salinidad.

Received: 26-12-2020          Accepted: 17-02-2021   Published: 24-02-2021

                                                                                                          INTRODUCTION
Because of the continuous production and supply of                     are grown under conditions that are not suitable for
tropical fruits, it is possible to increase cultivation                maximum development, with yields that are much
and export (Blancke, 2016). This opens a world of                      lower than the maximum recorded for the species as
possibilities for improving local and global consump-                  a consequence of suboptimal environmental condi-
tion with healthy foods (Viera et al., 2019). Most of                  tions (Raza et al., 2020). Thus, ecophysiology is close-
the so-called “exotic fruits” are important functional                 ly related to environmental conditions that affect
foods (Moreno et al., 2014; Campos et al., 2018) and                   plants when conditions are not optimal for growth
are highly valued, not only in tropical and sutropical                 and development. Very small changes in abiotic fac-
countries but also by consumers in higher latitudes                    tors, far from the optimum for the species, can mani-
(Ramadan, 2011). This trend has greatly benefited                      fest as stress for plants and have considerable effects
Andean countries with increasing export volumes                        on production (Jalil and Ansari, 2020).
since the beginning of the 21st century (Moreno-Mi-
randa et al., 2019).                                                   Climatic factors, such as temperature, relative hu-
                                                                       midity, vapor pressure deficit, solar radiation, rain
Plants require an abiotic environment to develop (Jalil                and wind, in addition to altitude, affect the ecophysi-
and Ansari, 2020). Thus, ecophysiological studies are                  ology of cultivated plants the most (Restrepo-Díaz
of utmost importance for the physical and biotic                       and Sánchez-Reinoso, 2020). At low latitudes, the
factors of the environment in terms of physiologi-                     inner tropics lack pronounced temperature seasons,
cal processes in plants and interaction mechanisms                     similar to temperate climate zones; consequently, the
that affect growth and development (Lambers et al.,                    rainy and dry seasons determine to which the plant
2008; Fischer et al., 2016). Environmental conditions                  physiology reacts and adapts (Fischer and Parra-
must be as close to optimal as possible for a planta-                  Coronado, 2020).
tion to achieve the highest crop production and qual-
ity, which are determined by the genetic potential                     Several abiotic factors always act simultaneously on
(Pérez and Melgarejo, 2015). Most of the time, plants                  plants, with little research (Raza et al., 2020); unlike

Rev. Colomb. Cienc. Hortic.
ECOPHYSIOLOGICAL ASPECTS OF GUAVA                     3

studies in controlled environments that allow only          has been introduced in many low-latitude countries
one or very few factors to vary, but which do not           (Singh et al., 2019). It has its highest production in
agree with the multidimensionality (Mittler, 2006)          India, Brazil and Mexico (Mishra et al., 2014) and on
in which all factors occur under field conditions. For      the continents South America, Asia and Australia
this reason, more ecophysiological studies on crops         (Singh et al., 2019). It is important for the domestic
are required because climate change requires the re-        economy of more than 60 countries in the tropics
evaluation of previous results (Restrepo-Díaz and           (Bandera and Pérez, 2015).
Sánchez-Reinoso, 2020).
                                                            The popularity of guava arises from its availability
According to Marengo et al. (2011), climate change          throughout the year, affordable price, high nutri-
will not affect low areas in the tropics as much. How-      tional and medicinal contents, aptitude for transport
ever, the Andean areas will increase rainfall by 20 to      and handling, and consumer preference (Methela et
25%. This phenomenon will consequently accelerate           al., 2019).
events such as “La Niña”, causing prolonged flood-
ing, accompanied by less solar radiation (Ramírez           In Colombia, Agronet (2020) reported a production
and Kallarackal, 2018; Sánchez-Reinoso et al., 2019;        of 80,814.7 t for 2018, harvested on 7,628.2 ha, main-
Arteaga and Burbano, 2018). Furthermore, climate            ly in the departments Boyaca (2,351.0 ha), Santander
change increases warming more in higher regions of          (1,790.6 ha), Tolima (841.0 ha) and Valle del Cauca
the tropics than in valleys (Marengo et al., 2011).         (693.6 ha). 69% is for fresh consumption, and the
                                                            rest is used for making sweets, known as bocadillo
Shukla et al. (2019) classified fruits and vegetables       (López-Santos et al., 2017).
within the species most affected by climate change,
where yield and quality tend to decrease as warm-           The guava plant belongs to the myrtaceae family,
ing increases, mainly in tropical and subtropical
                                                            known for its botanical richness and very high agro-
areas. Also, Haokip et al. (2020) attributed the fact
                                                            industrial potential with 121 genera and 5,800 spe-
that fruit trees have a higher incidence of physiologi-
                                                            cies of aromatic fruits, classified by Farias et al. (2020)
cal disorders, problems in pollination and changes in
                                                            as one of the more important commercial families
phenology to climate change. In addition, there are
                                                            globally.
many uncertainties in terms of the impacts of pests
and diseases in a changing climate (Tito et al., 2018),
                                                            Some 150 species belong to the genus Psidium (Ligar-
which influence fruit quality and food security.
                                                            reto, 2012), and most of the cultivars are P. guajava.
However, Devenish and Gianella (2012) and Raza et           However, there are other species of Psidium known as
al. (2020) stated that atmospheric warming can in-          P. cattleianum, P. molle, P. guineense, P. friedrichsthalia-
crease fruit production for trees in a given site. In ad-   num, and P. montanum, among others (Rai and Jaiswal,
dition, in the Andean region, there are suitable plots at   2020). P. guajava is the most cultivated in the world
higher altitudes that have the optimum temperature          (Aguilera-Arango et al., 2020). Most of the species of
of this crop (Tito et al., 2018) and optimal physiologi-    the genus Psidium are native fruit trees from the tro-
cal or ecological conditions. Likewise, DaMatta et al.      pical and subtropical Americas (Fischer et al., 2012).
(2010) concluded that C3 plants, which are almost
all fruit trees, could produce higher yields as the re-     The fruits have high levels of ascorbic acid (0.6-6.0
sult of increased atmospheric CO2 and use less water        g kg-1 edible fruit), vitamin A, calcium, phosphorus,
if no other stress conditions arise because of altered      potassium and dietary fiber (Paull and Duarte, 2012;
regional patterns of precipitation and temperature.         Prado et al., 2017). In addition, they are used because
                                                            of their antimicrobial, anti-inflammatory, antigeno-
Guava (Psidium guajava L.) is an evergreen tree that        toxic, and hepaprotective properties and for the treat-
is native to Mesoamerica and South America (Solarte         ment of diabetes and diarrhea (Pérez-Gutiérrez et al.,
et al., 2014), possibly from Mexico to Peru (Menzel,        2008). Its consumption reduces triglycerides, choles-
1985) or from Central America and southern Mexico           terol and blood pressure (Singh, 2007). Its flavor is
(Blancke, 2016). Paull and Duarte (2012) and Bandera        bittersweet, combined with a pleasant aroma, and it
and Pérez (2015) observed the origin as “simply” be-        is consumed fresh or processed into products such as
ing the tropical region of the Americas. Thanks to its      freshly cut salads, juice, nectar, cake, puree, concen-
ability to grow in tropical and subtropical climates, it    trates, jam, and gelatin, among others (Singh, 2011).

                                                                                                       Vol. 15 - No. 2 - 2021
4              FISCHER / MELGAREJO

Paull and Duarte (2012) characterized guava as a          fruiting occur continuously throughout the year in
shrub, which, under good humidity conditions, can         tropical and mild subtropical climates.
grow up to 6 to 9 m in height, forming trunks up
to 30 cm or more in diameter. By pruning and train-
ing, only one stem is developed, with a height of up      Temperature
to 3 or 4 m with flexible branches and young square       According to Gómez and Rebolledo-Podleski (2006)
twigs that become rounded with age. Yadava (1996)         and Salazar et al. (2006), the temperature range for the
described the guava tree as having undemanding            cultivation of guava is between 18 and 28ºC and be-
growth with a symmetrical and pyramidal shape. It
                                                          tween 15 and 30ºC, respectively; while Paull and Du-
forms bisexual flowers that are 2.5-3.0 cm in diam-
                                                          arte (2012) concentrated this range from 23 to 28ºC
eter, with autogamous reproduction, considerable
                                                          for optimal tree performance. Temperatures lower
self-pollination (60-75%) and 35% cross-pollination
                                                          and higher than this range reduce fruit setting, and
(Fischer et al., 2012; Menzel, 1985).
                                                          very low night temperatures (5-7ºC) paralyze growth
                                                          and turn leaves purple (Nakasone and Paull, 1998). If
The fruit contains many seeds and is botanically a
                                                          temperatures drop to 3ºC, the fruit no longer ripens
berry with spherical ovoid or pyriform shapes de-
                                                          (Insuasty et al., 2007). Sentelhas et al. (1996) found
pending on the variety, with diameters that vary
                                                          the lethal temperature for guava was -4ºC and rated
between 2.5 and 10.0 cm, skin colors between light
                                                          it as not very tolerant to low temperatures. Paull and
green and yellow and white to red pulps (Parra-
                                                          Duarte (2012) reported that prolonged low tempera-
Coronado, 2014). There are about 400 varieties in the
                                                          tures of -2ºC burn young plants.
world (Bandera and Pérez, 2015). Paull and Duarte
(2012) reported that this short-cycle fruit tree begins   Ferreira et al. (2019) estimated the cardinal develop-
to produce one year after planting, with maximum          ment temperatures for guava seedlings, applying 12
production at three or four years.                        different models with methodologies based on the
                                                          standard deviation of the accumulated degree days of
Because these promising crops have great potential        growth, calculating the base (minimum) temperature
for producers and international markets, ecophysi-        as 10.9ºC, the optimum as 17.3ºC and the maximum
ological impacts on quality and production charac-        as 51.2ºC. There is little information on cardinal
teristics (Mayorga et al., 2020) must be elucidated for   temperatures in fruit species, which are very useful
the development of guava crops. Therefore, the ob-        for studies on adaptation to different microclimates
jective of this review was to reveal climatic demands     (Souza and Martins, 2014).
and their effects on the physiology of plants, provid-
ing the basis of processes that have taken place for      Fruits react to unfavorable conditions for their devel-
the adaptation and diffusion of the species and useful    opment and quality. It has been observed that, at ele-
information for management and breeding programs.         vated temperatures, they are more aqueous, with low
                                                          sugar and ascorbic acid contents (Souza et al., 2010).
                                                          In addition, under conditions of high temperatures
                                                          and humidity during the development of the fruit,
ECOPHYSIOLOGICAL FACTORS AND THEIR
                                                          they become very susceptible to attacks by fruit flies
INFLUENCE ON GUAYABA                                      (Haokip et al., 2020).

There are diverse semi-wild and commercial forms,         Low temperatures, such as during the winter months
with very diverse morphological and nutritional           in the subtropics, make commercial production very
characteristics (Solarte et al., 2014); however, plant    difficult, increasing the time of fruit development to
breeding programs must develop cultivars with in-         about 220 days (Paull and Duarte, 2012), and, if the
creasingly superior fruit qualities and resistance to     cold season is also dry, these combined stress factors
abiotic and biotic stress (Thaipong and Boonprakob,       lead to natural defoliation and flowering will begin
2005). The guava tree blooms and produces at differ-      as soon as temperatures rise and rain induces a new
ent times depending on the site, climatic characteris-    flow of growth and fruit set (Nakasone and Paull,
tics, soil and crop management (Bandera and Pérez,        1998). Haokip et al. (2020) stated that flower drop
2015), as well as the genotype and the climatic condi-    occurs in guava when low temperatures prevail dur-
tions, which affect the growth cycle (Salazar et al.,     ing flowering. Floral opening depends on the daytime
2006). Singh (2011) confirmed that flowering and          temperature (Bandera and Pérez, 2015).

Rev. Colomb. Cienc. Hortic.
ECOPHYSIOLOGICAL ASPECTS OF GUAVA                     5

In subtropical areas, very high summer temperatures             allowed it to be distributed in the subtropics and nu-
can impair the content of sugars and organic acids in           merous countries of the world (Natale et al., 2008).
fruits, which are used in respiratory processes (Sol-
arte et al., 2014). In comparison, these authors found          Solarte et al. (2014) studied the effect of three alti-
an increase in ascorbic acid with increasing tempera-           tudes (1,570; 1,720 and 1,890 m a.s.l.) on fruit quality
ture and relative humidity at tropical altitudes be-            in four guava genotypes grown in a traditional semi-
tween 1,570 and 1,890 m a.s.l.                                  wild system in the Department of Santander, Colom-
                                                                bia with a bimodal rainfall regime (average rainfall/
Salazar et al. (2006) measured the phenological                 year of 1,780 mm and temperature of 20°C). In this
stages of guava by also calculating the degree days             study, the environmental factors that were associated
of growth, adding the differences between the mean              with altitude and resulted in differences in fruit quali-
daytime temperatures and the base temperature of                ty between the genotypes included the vapor pressure
12.0ºC, which was recorded when the development                 deficit (VPD), the maximum photosynthetic photon
of the flower bud began, comparing the development              flux density (PPFDmax) and the temperature differ-
of the plant between the different years and geo-               ence between day and night (∆T°). It was observed
graphical sites.                                                that, at low altitudes (with higher temperature and
                                                                solar radiation), the fruits had a higher fresh weight
                                                                and changed faster from green to yellow; while, in
Radiation                                                       general, the content of organic acids increased with
                                                                increasing altitude. Likewise, the higher altitude con-
Paull and Duarte (2012) found that a greater num-
                                                                ditions promoted efficiency in the accumulation of
ber of hours of sunlight leads to greater growth of
                                                                the monosaccharides fructose and glucose, but only
the branches. Also, the concentration of ascorbic acid
increases with increasing light intensities; however,           in two of the four genotypes. Solarte et al. (2014)
Solarte et al. (2014) recorded a decrease in this acid          stated that there was a genotype×environment in-
with radiation >2,000 µmol photons m-2 s-1.                     teraction effect on all variables because not all geno-
                                                                types reacted uniformly.

Altitude                                                        Musyarofah et al. (2020) evaluated ‘Kristal’ guavas
                                                                from low (200 m a.s.l.) and middle (550 m a.s.l.) al-
According to Solarte et al. (2014), the ecophysiologi-          titudes in Indonesia and found that fruits from the
cal effect of altitude on the guava plant depends               low elevation were heavier and bigger, with a higher
mostly on the variety but the concept of multidi-               vitamin E-content than in those grown in the middle
mensionality of Mittler (2006) must be considered               altitude farm, with fruits that were crispier and not
because increasing altitudes decrease the temperature           as soft as the low elevation ones.
(about 0.6ºC/100 m), the partial pressure of the air
(O2, CO2 and N2) and the relative humidity, while
radiation (visible, UV and infrared), rain (from 1,300-         Water
1,500 m a.s.l.) and wind increase (Fischer and Orduz-
Rodríguez, 2012).                                               Water is essential for all reproductive phases of guava
                                                                (Fischer et al., 2012). In a high Andean phenological
However, since guava is a native species of the trop-           study on the agrometeorological influences on the
ics, adaptation to these ranges of altitude (Tab. 1) has        reproductive phase of plants, Mendoza et al. (2017)

 Table 1. Recommended altitudes for growing guava.

            Altitude (m a.s.l.)                         Annotation                                   Author
                0 – 2,300                                 Ecuador                                Morton (1987)
                0 – 2,000                   Grows in a wide range of altitudes                Solarte et al. (2014)
                0 – 1,800                                Venezuela                               Hoyos (1989)
                0 – 1,700                    This range favors the distribution               Natale et al. (2008)
                0 – 1,500                            In frost-free places                    Paull and Duarte (2012)

                                                                                                              Vol. 15 - No. 2 - 2021
6              FISCHER / MELGAREJO

found that rain is the climatic factor that most pro-       suggested drip irrigation. In larger plantations that re-
motes this reproductive phase (73.4%), when com-            ceive irrigation by sectors, they recommend sprinkler
pared to temperature (19.3%), followed by solar             or microjet systems that, additionally, provide the
radiation or the photoperiod (3.2%).                        nutrients quickly to the plant. Likewise, to guarantee
                                                            the economic sustainability of a crop, Aguilar-Arango
Salazar et al. (2006) reported that guava trees need        et al. (2020) suggested the application of irrigation,
a water supply of about 1,000 to 2,000 m3 ha-1 year-1,      modification of the pruning season, and scheduling
but Aguilera-Arango et al. (2020), in areas with a bi-      the main harvest taking into account the fact that
modal system, annual rainfall of 800 to 1,300 mm            pruned plant needs enough water for the develop-
that is well-distributed, saw good development and          ment of new shoots.
production of the crop. On the other hand, Natale
et al. (2008) stated that the ideal range of rainfall is    Crane et al. (2019) classified the guava tree as tolerant
1,000 to 1,600 mm, well-distributed throughout the          to waterlogging, while Morton (1987) categorized
year, but it should not be less than 600 mm year-1. In      it as moderately tolerant to waterlogging. Kongsri
turn, because of the relative tolerance of guava to wa-     et al. (2020) observed that guava trees propagated
terlogging (Crane et al., 2019), Hoyos (1989) reported      by seedlings were more tolerant than those propa-
that these plants can be grown in regions with rain-        gated by shoot layering under flooding conditions.
fall between 1,000 and 3,000 mm per year; however,          Solarte et al. (2010) reported that prolonged rainfall
the same author reported that periods of excessive          can cause alterations in the normal production cycle
rains during the development of the fruit can cause         in the Suarez river basin (Department of Santander,
cracking and harvest losses.                                Colombia), which shorten harvest times. On the
                                                            other hand, these authors observed an increase in
Excessive rains during fruit development make them
                                                            the foliar anthocyanin pigments that decrease the
more watery, with less firmness and reduced con-
                                                            photosynthetic capacity and thus the production of
tents of sugars, titratable acidity and ascorbic acid.
                                                            guava. In addition, very humid conditions increase
According to Souza et al. (2010), the values ​​varied
                                                            diseases and pests (Singh, 2011), causing the abor-
depending on the precipitation volumes and the rip-
                                                            tion of a large number of fruits (Solarte et al., 2010).
ening stages of the fruits. Menzel (1985) confirmed
                                                            Likewise, environments with prolonged high rela-
that excess water during the fruiting period increases
                                                            tive humidity damage the quality of guavas (Fischer
the cracking and fall of fruits, similar to those of cape
                                                            et al., 2012).
gooseberry (Fischer and Melgarejo, 2020). Sharma et
al. (2020) reported that, in India, many crops in the
                                                            Taiwo et al. (2020) pointed out that drought is the
summer rainy season are discarded because of poor
                                                            most prevalent abiotic stress in the world, which
commercial quality and found that bagging fruits
                                                            limits the productivity of plantations. Therefore, in
with polypropylene non-woven bags is very benefi-
                                                            the dry tropics, the flowering of the guava is high-
cial for controlling pests and diseases and improving
                                                            ly influenced by the availability of water (Paull and
the harvest quality during the rainy season.
                                                            Duarte, 2012). Interestingly, guava is not only classi-
The favorable contribution of water in guava is not         fied as tolerant to waterlogging but also moderately
only essential for full vegetative growth but also in       tolerant to droughts (Crane et al., 2019). Alix et al.
the beginning of the reproductive phase, in flower-         (cited by Paull and Duarte, 2012) mentioned that this
ing, and in the setting and filling of fruits; therefore,   plant can withstand a dry period of about 6 months.
in nature, flowering begins with the rainy season           This tolerance is surprising because of the superficial
(Fischer et al., 2012; Paull and Duarte, 2012). The         root system (Menzel, 1985). This tolerance is fa-
sprouting of the terminal branches that will carry          vored by the large number and extension of the roots
the flowers requires an optimal supply of humidity          that exceed the diameter of the crown (Bandera and
(Mata and Rodríguez, 2000). These same authors re-          Pérez, 2015). As previously mentioned, dry seasons
ported that a minimum rainfall of 127 to 178 mm             combined with cold temperatures usually induce
per month is required in Hawaii and that irrigation         defoliation of the tree, which recovers fully if these
is applied to advance flowering, which also allows          conditions change (Nakasone and Paull, 1998). Fur-
scheduling the harvest (Aguilera-Arango et al., 2020).      thermore, as Fischer et al. (2012) mentioned, low hu-
                                                            mid periods promote flower induction; while, on the
To replace the evaporated water in a guava planta-          contrary, dry conditions can induce the abortion of
tion (25-50 mm/week), Nakasone and Paull (1998)             already formed flowers.

Rev. Colomb. Cienc. Hortic.
ECOPHYSIOLOGICAL ASPECTS OF GUAVA                  7

Irrigation is necessary in regions with long dry sea-      nitrogen fertilizers as alternatives for agricultural
sons, which is why Paull and Duarte (2012) empha-          production in these regions (Souza et al., 2017).
sized that the ideal pattern for guava is alternating
conditions of dry and humid seasons since drought          Salinity, especially at levels greater than 1.8 dS m-1,
and low ambient humidity during flowering can seri-        affects the emergence of seedlings, as well as growth
ously reduce fruit set and cause abscission of recently    and biomass accumulation in guava. The cultivar
set fruit. Likewise, these authors stated that, when       ‘Crioula’ is more tolerant to salinity than ‘Paluma’
trees that suffer from water stress abort the fruits of    and ‘Ogawa’ and is recommended as a rootstock (Sá
1-2 cm in diameter after intense irrigation or heavy       et al., 2016). Cavalcante et al. (2005) found that four
rain, these trees resume vegetative growth. In addi-       guava varieties with seedlings irrigated with an ECw
tion, low humidity during fruit filling reduces size       greater than 1.5 dS m-1, 180 d after sowing, did not
and causes a recollection of the pulp from the epider-     have the agronomic quality for transplanting; while
mis (Paull and Duarte, 2012).                              Ramírez et al. (2017) observed, in the germination,
                                                           longitudinal root and stem growth of the variety
As reported by Souza et al. (2010), the hydric condi-      ‘Criolla Roja’, a slight tolerance up to a concentration
tions in guava promote optimum fruit quality, and          of 2.5 dS m-1 of NaCl.Souza et al. (2020) found that P.
excess water during fruit filling reduces the soluble      cattleianum is not very suitable as a rootstock in saline
solids content; while the sugar concentration is fa-       areas because of its greater absorption of Na+, which
vored by water scarcity as a concentration effect          leads to high levels of Na+ in the leaves of the scion
(Mercado-Silva et al., 1998).                              and, thus, lower tolerance to saline stress.
In drier environments, Solarte et al. (2014) found that,   In the Paluma variety, increases in irrigation water
as the air vapor pressure deficit (DPV) and the maxi-      salinity from 0.3 dS m-1 produced reductions in sto-
mum photonic flux density (PPFDmax) increased, the         matal conductance, internal CO2 concentration, CO2
concentrations of citric and succinic acid decreased       assimilation rate, transpiration, and efficiency in the
in guava. The DPV and PPFDmax can influence the            instantaneous use of water, in addition to reducing
water status in fruits (Genard et al., 2009) and thus      the number of leaves and branches, the diameter of
reduce the transpiration and photosynthesis rates of       the stem and the absolute and relative growth rates
green fruits, affecting the primary metabolism of this     (Bezerra et al., 2018a). However, in this study, the
organ.                                                     growth of ‘Paluma’ was affected by increases in the
                                                           water salinity, and these plants could be irrigated
                                                           with water of up to 1.42 dS m-1, causing an accept-
Salinity
                                                           able reduction of 10% in growth variables.
Jalil and Ansari (2020) reported that salinity occurs
                                                           Cavalcante et al. (2010) managed to relieve the ef-
in areas with little rainfall and high temperatures,
which promote high transpiration rates and affect          fects of salinity with the application of liquid bovine
the normal development of plants because of the use        manure, but potassium applications failed to do so
of saline water for irrigation, where excess sodium        (Bonifácio et al., 2018). Applications of N above 70%
and chlorine ions cause toxicity and hinder the ab-        of the recommended dose (378.7 mg N dm-3 soil) did
sorption of essential elements and water from the          not mitigate the detrimental effect of saline stress on
soil.                                                      plants (Bezerra et al., 2018b).

The guava tree shows moderate tolerance to salinity
(Morton, 1987), and applications of calcium nitrate,       Winds
Ca(NO3)2 of 10 mM, alleviatereductions in growth
ofseedlings induced by NaCl through an increase in         Paull and Duarte (2012) recommended windbreaks
the concentration of foliar chlorophyll and higher         for guava, especially for high-quality dessert-type
photosynthetic rates (Ebert et al., 2002).                 fruits produced for the fresh market. In addition,
                                                           these authors reported that plants grafted on clon-
Many studies on salinity in guava were carried out         al rootstocks are very susceptible to wind speeds
in Brazil, especially in arid and semi-arid areas, where   between 65 and 80 km h-1 during the first three years
the scarcity of good quality water and the occurrence      of cultivation; while trees exposed to winds between
of low fertility soils are limiting factors in irrigated   16 and 32 km h-1 have branches that gradually de-
agriculture, which led to the use of salt water and        velop out of the direction of the wind.

                                                                                                   Vol. 15 - No. 2 - 2021
8              FISCHER / MELGAREJO

On the contrary, Bandera and Pérez (2015) reported              Estado actual de la investigación para el cultivo de
that, despite the fact that the root system of guava is         guayaba en Colombia. Agron. Mesoamer. 31(3), 845-
superficial, it resists strong winds and storms thanks          860. Doi: 10.15517/am.v31i3.40207
to the extension and number of large roots that ex-        Arteaga, L. and J. Burbano. 2018. Efectos del cambio climá-
ceed the width of the canopy of the crown, allowing             tico: Una mirada al Campo. Rev. Cienc. Agríc. 35(2),
this species to develop in a large number of different          79-91. Doi: 10.22267/rcia.183502.93
soils.                                                     Bandera, E. and L. Pérez. 2015. Mejoramiento genético de
                                                               guayabo (Psidium guajava L.). Cult. Trop. 36(Sp. Tiss.),
                                                               96-110.
CONCLUSIONS                                                Bezerra, I.L., R.G. Nobre, H.R. Gheyi, G.S. Lima, and J.L.
                                                                Barbosa. 2018a. Physiological indices and growth of
Guava, thanks to its ability to grow in tropical and            ‘Paluma’ guava under saline water irrigation and ni-
subtropical climates, has been introduced to many               trogen fertigation. Rev. Caatinga 31(4), 808-816. Doi:
low-latitude countries.                                         10.1590/1983-21252018v31n402rc
                                                           Bezerra, I.L., R.G. Nobre, H.R. Gheyi, L.P. Souza, F.W.A.
This crop grows in temperature ranges between 15                Pinheiro, and G.S. Lima. 2018b. Morphophysiology of
and 30°C. Lower and higher temperatures reduce                  guava under saline water irrigation and nitrogen ferti-
fruit set. Low temperatures make production diffi-              lization. Rev. Bras. Eng. Agríc. Ambient. 22(1), 32-37.
cult, causing flower drop or extending the fruit de-            Doi: 10.1590/1807-1929/agriambi.v22n1p32-37
velopment phase. However, guava adapts well up             Blancke, R. 2016. Tropical fruits and other edible plants of
to altitudes of 2,000 m a.s.l. Colombia has a high              the world. Cornell University Press, Ithaca and Lon-
genotype×environment interaction in the produc-                 don. Doi: 10.7591/9781501704284
tion characteristics and fruit quality with respect to
                                                           Bonifácio, B.F., R.G. Nobre, A.S. Sousa, E.M. Gomes, E.M.
the elevation of the orchard.                                   Silva, and L.P. Sousa. 2018. Efeitos da adubação po-
                                                                tássica e irrigação com águas salinas no crescimento
An important ecophysiological factor in guava is                de porta-enxerto de goiabeira. Rev. Ciênc. Agr. 41(4),
water since crops require between 1,000 to 2,000                971-980. Doi: 10.19084/RCA18119
mm year-1 and endure waterlogging for several days;
                                                           Campos, D., R. Chirinos, L. Gálvez, and R. Pedreschi. 2018.
however, a lot of precipitation and environmental
                                                              Bioactive potential of Andean fruits, seeds, and tu-
humidity decrease fruit quality considerably. On the          bers. pp. 287-343. In: Toldra, F. (ed.). Advances in food
other hand, since this tree is classified as moderately       and nutrition research. Vol. 84. Elsevier, Cambridge,
drought tolerant, this adversity greatly affects flow-        MA. Doi: 10.1016/bs.afnr.2017.12.005
ering and fruit set. In addition, since it has moderate
                                                           Cavalcante, L.F., Í.H.L Cavalcante, K.S.N. Pereira, F.A. Oli-
tolerance to salinity, it supports irrigation water with
                                                               veira, S.C Gondim, and F.A.R. Araújo. 2005. Germina-
up to an EC of 1.5 to 1.8 dS m-1 depending on the              tion and initial growth of guava plants irrigated with
variety.                                                       saline wáter. Rev. Bras. Eng. Agríc. Ambient. 9(4), 515-
                                                               519. Doi: 10.1590/S1415-43662005000400012
Guava should be researched under conditions of el-
evated CO2 concentrations, along with interactions         Cavalcante, L.F., M.S. Vieira, A.F. Santos, W.M. Oliveira,
                                                               and J.A.M. Nascimento. 2010. Água salina e esterco
with nitrogen fertilization.
                                                               bovino líquido na formação de mudas de goiabeira
                                                               cultivar Paluma. Rev. Bras. Frutic. 32(1), 251-261. Doi:
Conflict of interests: The manuscript was prepared
                                                               10.1590/S0100-29452010005000037
and reviewed with the participation of the authors,
who declare that there exists no conflict of interest      Crane, J.H., C.F. Balerdi, and B. Schaffer. 2019. Managing
that puts at risk the validity of the presented results.       your tropical fruit grove under changing water table
                                                               levels. Doc. HS957. Horticultural Sciences Depart-
                                                               ment, UF/IFAS Extension, Gainesville, FL.
                                                           DaMatta, F.M., A. Grandis, B.C. Arenque, and M.S. Bucke-
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